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Solanaceae) Complex in Costa Rica

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Solanaceae) Complex in Costa Rica BIOTROPICA 32(1): 70–79 2000 Insights into the Witheringia solanacea (Solanaceae) Complex in Costa Rica. I. Breeding Systems and Crossing Studies1 Lynn Bohs2 Department of Botany, Duke University, Durham, North Carolina 27708, U.S.A. ABSTRACT The Witheringia solanacea complex consists of three species, W. asterotricha, W. meiantha, and W. solanacea, native to Central and South America. The three taxa are morphologically similar, and their distinctions and relationships have been the subject of taxonomic controversy. To investigate breeding systems and potential for hybridization among the taxa of the complex, two Costa Rican accessions per species were used in a crossing program. All plants were self- incompatible except for one accession of W. solanacea. Hybrid plants resulted from all crosses among accessions of W. asterotricha and W. solanacea. Most crosses were unsuccessful using W. meiantha in combination with either of the other two taxa. It is suggested that W. meiantha and W. solanacea be recognized as separate taxa, but that W. asterotricha be considered a synonym of W. solanacea. RESUMEN El complejo Witheringia solanacea consiste de tres especies de Centro y Surame´rica, W. asterotricha, W. meiantha, y W. solanacea. Las tres especies son morfolo´gicamente parecidas entre sı´ y han sido fuente de muchas controversias tax- ono´micas. Para investigar el sistema reproductivo y el potencial para hibridizacio´n entre las especies del complejo, se utilizaron seis colecciones de Costa Rica (dos por especie) en un programa de cruzamiento en invernadero. Todas las plantas eran auto-incompatibles salvo una coleccio´n de W. solanacea. Se formaron hı´bridos en todas las combinaciones entre las colecciones de W. asterotricha y W. solanacea. La mayorı´a de intentos entre W. meiantha y las otras dos especies fallaron. Se sugiera un esquema taxono´mico para el complejo que reconoce dos especies, W. meiantha y W. solanacea, con W. asterotricha como sino´nimo de W. solanacea. Key words: breeding systems; Costa Rica; crossing studies; self-incompatibility; Solanaceae; Witheringia. THE GENUS WITHERINGIA L’H ER.(SOLANACEAE) IN- ingia may be more closely related to the physaloid CLUDES ca 20 species of Neotropical herbs, shrubs, genera [Physalis L., Margaranthus Schlecht., Leu- and small trees. The center of diversity for the ge- cophysalis Rydb., Chamaesaracha (A. Gray) Benth.] nus is Costa Rica, where over half the species occur. and that the genus may not be monophyletic as Witheringia has been placed in subfamily Solan- currently circumscribed (Olmstead et al. 2000; oideae, tribe Solaneae, by virtue of its flattened Bohs, pers. obs.). seeds with curved embryos, abundant endosperm, The genus Witheringia is distributed from basal filament insertion, and valvate corolla aesti- Mexico to Bolivia, with the majority of species oc- vation (Hunziker 1979, D’Arcy 1991). Witheringia curring in Mexico, Costa Rica, and Panama. Hun- has been allied with the genera Acnistus Schott, ziker’s (1969) revision treated 15 taxa, including 3 Athenaea Sendtn., Aureliana Sendtn., Brachistus species of section Brachistus. He pointed out that Miers, Capsicum L., Cuatresia Hunz., Dunalia 3 species, W. asterotricha (Standl.) Hunz., W. H.B.K., Iochroma Benth., Saracha Ruiz & Pav., and meiantha (Donn. Sm.) Hunz., and W. solanacea Vassobia Rusby in traditional classification schemes L’Her., form a tightly related group termed the W. (Hunziker 1984, 1987) and often has been consid- solanacea complex. All are herbs or shrubs with ered as being closely related to Capsicum because truncate calyces and a ring of hairs inside the co- of similarities in flower and fruit characters. Recent rolla tube, and with corolla lobes much longer than molecular studies, however, indicate that Wither- the tube. These species have very short (Ͻ5 mm) peduncles or lack them altogether, so that the flow- 1 Received 15 January 1998; revision accepted 7 Novem- ers are arranged in fascicles in branch forks or op- ber 1998. posite the paired leaves. Witheringia solanacea has 2 Current address: Department of Biology, University of Utah, Salt Lake City, Utah 84112, U.S.A.; e-mail: the broadest geographical distribution, ranging [email protected]. from southern Mexico through Central America 70 Crossing Studies in Witheringia 71 and the Antilles to South America. It is an extreme- ductive interaction among the taxa is at least the- ly variable species, but is distinguished by its pu- oretically possible. bescence of unbranched or sparsely branched hairs In addition to the crossing results, a brief key and often tetramerous flowers. Hunziker (1969) to the three entities is included at the conclusion listed one variety, W. solanacea var. silvigaudens of the paper. A complete systematic treatment of (Standl. & Williams) Hunz., endemic to Dept. the genus Witheringia is being prepared by M. Sou- Moraza´n, Honduras. This variety is glabrous, has sa-Pen˜a of the University of Connecticut. pentamerous corollas, and longer, narrower leaves than typical W. solanacea. Witheringia asterotricha MORPHOLOGY is closely related to W. solanacea, and according to Hunziker (1969), probably should be considered a The three taxa of the complex are all erect herbs, variety of the latter species. Witheringia asterotricha, weakly woody shrubs, or small trees reaching sev- thus far known only from Costa Rica and Panama, eral meters in height. The leaves are entire, elliptic, is morphologically distinguishable from W. solana- unequal in size, and paired at the nodes (Fig. 1A). cea by its dense pubescence of dendritically All three taxa lack obvious peduncles, and as a re- branched hairs and pentamerous flowers. Wither- sult, flowers are clustered in axillary fascicles (Fig. ingia meiantha ranges from southern Mexico to 1). The corollas are stellate and light yellowish to western Panama. Plants of W. meiantha are gla- cream in color, with those of W. meiantha smaller brous, with small pentamerous flowers and fewer- and deeper yellow than those of W. asterotricha and flowered inflorescences than W. solanacea or W. as- W. solanacea. The latter two taxa often have green- terotricha. ish maculations in the corolla throat, and the co- Unlike Hunziker (1969), D’Arcy (1973) rec- rolla lobes are longer and less reflexed than in W. ognized only two species in the complex. He main- meiantha. The tapered anthers are connivent tained W. asterotricha as distinct, but indicated that around the style, and begin to dehisce by large ter- minal pores that eventually open into longitudinal hybrids with W. solanacea may occur when the two slits (Fig. 1B). After pollination, the corolla and species come into contact. D’Arcy (1973) consid- attached stamens wither and fall, leaving only the ered W. meiantha a synonym of W. solanacea, com- calyx and gynoecium. As fruits develop, the pen- menting that the reduced pubescence and slightly dulous flowering pedicels curve upward until the larger calyces characterizing W. meiantha were like- ripe fruits are held erect (Fig. 1A, C, D). At ma- ly minor morphological variants of the widespread turity, all three taxa have red, shiny, juicy, many- W. solanacea. seeded fruits. Witheringia meiantha has glabrous Nee (1986), treating only the taxa from Vera- fruits, those of W. solanacea are glabrous to very cruz, Mexico, recognized W. meiantha as distinct sparsely pubescent, and those of W. asterotricha are from W. solanacea and considered W. solanacea var. moderately to rather densely pubescent. Fruit color silvigaudens a synonym of W. meiantha. According changes from green to orange–red very quickly, of- to Nee (1986), W. meiantha can be distinguished ten in a single day. These attributes are character- from W. solanacea by its larger calyx and seeds, pen- istic of dispersal by birds, or ornithochory (van der tamerous flowers, glabrous leaves, and few-flowered Pijl 1982). Bird dispersal of Witheringia fruits has inflorescences. been documented at La Selva and Monteverde, To gain further insight into the relationships of Costa Rica, by Wheelwright et al. (1984), Murray these taxa, I carried out a crossing study in the (1987, 1988), Loiselle and Blake (1990), Blake and greenhouse to determine if genetic isolating mech- Loiselle (1992), and Murray et al. (1994). anisms were operating among members of the W. solanacea complex. It was also of interest to ex- amine breeding systems of the three taxa, as no STUDY SITES AND HABITATS information of this kind is available for Withering- One accession of W. asterotricha and both acces- ia, and the distribution of self-incompatibility (SI) sions of W. meiantha used in the crossing study among genera of the Solaneae is not well known. were collected at La Selva Biological Station near Greenhouse plants were grown from seed collec- the confluence of the Sarapiquı´ and Puerto Viejo tions (accessions) of three sites in Costa Rica. Costa Rivers in Heredia Province (Table 1). Elevations at Rica, and perhaps Panama, are the most likely areas the station range from ca 35 to 140 m. La Selva of sympatry or near sympatry for the three species, has an annual mean temperature of 26ЊCandan and therefore represent localities in which repro- annual mean rainfall of ca 4000 mm. Although 72 Bohs FIGURE 1. A. Witheringia asterotricha branch and inflorescence with flowers and young fruits. B. Flower of W. solanacea. C. Fruits of W. asterotricha. D. Flowers and fruits of W. meiantha. rain falls at La Selva in all months of the year, there (Route 9) near the Catarata de La Paz at an elevation is a relatively wet season from late April to the of ca 1350 to 1400 m. Climatic data are not available beginning of August, with a smaller wet period for the site, but the average temperature is lower than from November through January. Detailed descrip- at La Selva, and the annual rainfall is probably greater. tions of the site can be found in McDade et al. One accession of W. solanacea was collected here at a (1994).
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